A reassessment of the Neanderthal teeth from Taddeo cave (southern Italy)

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Journal of Human Evolution 61 (2011) 377e387

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Journal of Human Evolution

journal homepage: www.elsevier .com/locate/ jhevol

A reassessment of the Neanderthal teeth from Taddeo cave (southern Italy)

Stefano Benazzi a,*, Bence Viola a,b,c, Ottmar Kullmer d, Luca Fiorenza d,e, Katerina Harvati f, Tobias Paul a,Giorgio Gruppioni g, Gerhard W. Weber a, Francesco Mallegni h

aDepartment of Anthropology, University of Vienna, Althanstraße 14, 1090 Vienna, AustriabDepartment of Human Evolution, Max Planck Institute of Evolutionary Anthropology, Detuscher Platz 6, D-04103 Leipzig, GermanycDepartment of Evolutionary Genetics, Max Planck Institute of Evolutionary Anthropology, Detuscher Platz 6, D-04103 Leipzig, GermanydDepartment of Palaeoanthropology and Messel Research, Senckenberg Research Institute, Senckenberganlage 25, D-60325 Frankfurt am Main, GermanyeArchaeology & Palaeoanthropology, University of New England, Armidale NSW 2351, Australiaf Paleoanthropology, Department of Early Prehistory and Quaternary Ecology and Senckenberg Center for Human Evolution and Paleoecology, Rümelinstrasse 23,Tübingen 72070, GermanygDepartment of History and Method for the Conservation of Cultural Heritage, University of Bologna, Vicolo degli Ariani 1, 48100 Ravenna, ItalyhDepartment of Biology, University of Pisa, Via S. Maria 53, 56126 Pisa, Italy

a r t i c l e i n f o

Article history:Received 18 June 2010Accepted 5 May 2011

Keywords:Middle PaleolithicDental remainsMicro-Computed TomographyDental tissue proportionsVirtual reconstructionEnamel thickness

* Corresponding author.E-mail address: [email protected] (S. Be

0047-2484/$ e see front matter � 2011 Elsevier Ltd.doi:10.1016/j.jhevol.2011.05.001

a b s t r a c t

TheMiddle Paleolithic fossil human teeth fromTaddeo cave in southwestern Italywere discovered in 1967,but to date only scanty and partially incorrect information has been published about them. The teeth wererecovered in a reddish sandy layer fromthe cave’sfloor,which is attributedeither to an early phase ofWürmI (OIS 5c or 5d) or a transition phase between Würm I and Würm II (OIS 5a). In this paper, we presenta revisedmorphological description andmorphometric comparisons of the fourdental remains discovered.Apart from a classic morphometric comparison, we also provide a qualitative and quantitative assessmentof the internal morphology with the aid of micro-CT imaging. In addition, virtual restoration andmatchingof adjacent teeth were performed with 3D digital modeling and Computer-Aided Design techniques.Occlusal Fingerprint Analysis was also employed to help correctly identify each tooth.

While in the previous studies, Taddeo 1 was considered either an upper right canine or a lower rightcanine, in the present work it has been definitely identified as lower left canine. Taddeo 2 has beenreclassified as a right P4 instead of a right P3. Based on the occlusal and interproximal wear, we have alsoshown that Taddeo 2 and Taddeo 3 (right M1) belong to the same individual.

All of the teeth show characteristic Neanderthal features in crown morphology and fissure pattern.However, although Taddeo 4 shows morphological features typical of Neanderthal M1s, some morpho-metric results (large enamel thickness, low dentine volume) recall more modern humans than Nean-derthals. This result might suggest that, at least for lower first molars, the Neanderthal range of variationis large and still not clearly understood.

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Introduction

In 1967, during a systematic excavation of Taddeo cave (locatedon La Calanca beach, along the Marina di Camerota coast, Campa-nia, Italy), four human teeth were discovered in a layer withMousterian lithic industry (Vigliardi, 1968; see below). These teethwere described by Messeri (1975), who observed modern humanfeatures along with Neanderthal traits, suggesting that duringWürm I modern human populations lived on the Italian coasttogether with Pre-Neanderthals (Messeri, 1975; Messeri and Palma

nazzi).

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di Cesnola, 1976). Even though a new description of these remainswas not carried out, there is a general belief among paleoanthro-pologists that these teeth belong to Neanderthals, both in relationto the sediment layer from which they come from and based onsome typical Neanderthal features recognizable in the teeth.Consequently, without specifying their taxon but pointing out theirassociation with a Mousterian industry, these teeth were includedin the Catalog of Italian Fossil Human Remains (Giacobini andManzi, 2005) as follows: Taddeo 1¼ lower right canine (RC1);Taddeo 2¼ upper right third premolar (RP3); Taddeo 3¼ upperright first molar (RM1); Taddeo 4¼ lower right first molar (RM1).

There are several reasons that justify a reassessment of theseteeth. First, there are arguments that seem to challenge Messeri’sconclusions (Messeri, 1975; Messeri and Palma di Cesnola, 1976)

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S. Benazzi et al. / Journal of Human Evolution 61 (2011) 377e387378

about their taxonomic assignment. Second, recent observationscarried out by one of the authors (SB) led to dissenting conclusionsabout the classification of at least two specimens from Taddeo cave,namely, Taddeo 1 and Taddeo 2. Third, the last description wascompleted more than 30 years ago (Messeri, 1975), without theassistance of current technological advances that allow non-invasive access to the internal structure of teeth (i.e., to comparethe enamel-dentine junction e EDJ, or to quantify the enamelthickness), and before the systematic description of Neanderthaldental non-metric traits was undertaken (Bailey, 2002). Fourth,during this time period these fossil remains were seriouslydamaged (see below), underscoring the need for extensive inves-tigation and proper documentation of these specimens.

In this contribution, we present the first detailed morphologicaldescription andmorphometric comparison of the four human teethfrom Taddeo cave. In addition to morphological and morphometricanalyses of the external aspects of the teeth, we also providequalitative and quantitative information of the internal structureobtained by micro-Computed Tomography (mCT) investigation.Furthermore, based on 3D digital modeling and Computer-AidedDesign (CAD) techniques, a virtual restoration was performed forsome teeth to collect volumetric data, which we believe will beuseful for future comparative studies.

Taddeo cave (Fig. 1)

Taddeo cave consists of a small chamber with a cave floor ofelliptical shape (ca. 16 m� 10 m). The stratigraphic sequence of thedeposition is described as being simple, however, a drawing of thestratigraphic section is not available. The sequencewas described asconsisting of, from the bottom to the top, a layer of gray sandstone(the Tyrrhenian beach, which was the trampling surface during theuseof the cave), overlaid bya thin reddish sandy layer that is coveredby a surface speleothem of variable thickness (Vigliardi, 1968). Thereddish sandy layer yielded faunal remains, lithic artefacts, and thefour human teeth that we re-investigate here. The animal remainswere studied by E. Borzatti von Löwenstern (Palma di Cesnola,1996)and were attributed to several large mammal taxa: Hippopotamusamphibius, Dicerorhinus mercki, Cervus elaphus, Capreolus capreolus,Sus scrofa, Capra ibex and, in lesser quantity, remains of Bos pri-migenius and Equus caballus. The Hippopotamus remains indicatethat a permanent freshwater source such as a lake, river or a swamp,surrounded by trees and bushes, existed in the vicinity of Taddeocave. However, the horse and rhino remains also suggest a grasslandenvironment not far from the cave (Vigliardi, 1968; Martini et al.,1976; Palma di Cesnola, 1996, 2001).

Figure 1. Geographical position of Taddeo cave, whose entrance is located on LaCalanca beach (Marina di Camerota, Salerno, Italy).

The chronology of this layer has not been resolved. Since thesefaunal remains are indicative of a wet-warm climate, the layer hasbeen attributed to an early phase of the Würm I (OIS 5c or 5d)(Vigliardi, 1968). Nevertheless, this chronology was questioned byPalma di Cesnola (1996) on the basis of the lithic artifacts found inthe same layer. Only a few pieces were recovered, including fivesidescrapers, six retouched splinters, four denticulates and onenotched piece. Almost all of these were made on flat flakes usuallyslightly retouched, partly produced using a Levallois technique. Thelithic tools show characteristics usually found in Mousterianindustries (Vigliardi, 1968; Palma di Cesnola, 2001), but they do notcorrespond to the more ancient Mousterian found in the regionelsewhere. Accordingly, Palma di Cesnola (1996) suggested a phasebetween Würm I-II (OIS 5a, closer to OIS 4) as a more appropriategeological age for the assemblages. On the basis of the faunalassemblage, as well as of the small size of the lithic artefacts, anephemeral use of Taddeo cave has been hypothesized.

Methods

Traditional analysis methods

For each of the four dental specimens, both metric and non-metric data were collected. Terminology used for cusps, crestsand furrows follow Scott and Turner (1997). Non-metric characterswere recorded using the Arizona State University Dental Anthro-pology System (ASUDAS) (Turner et al., 1991), supplemented by thecharacters described by Bailey (2002, 2006). Occlusal wear stagewas assessed through visual examination and scored based on theattrition categories of Molnar (1971). Standard dental measure-ments (e.g., mesio-distal (MD) and buccolingual (BL) diameters)were taken with digital calipers. The MD crown diameter wasdefined as the maximum distance between the mesial and distalcrownwalls, aligning the caliper parallel to the occlusal surface. Forthe molars, we used the mesial face as a reference plane for posi-tioning the mesial caliper point, while the distal measurementpoint is consequently determined through closing the caliper(Martinón-Torres et al., 2008). For the canine and the premolar, theBL diameter was defined as the maximum width between thebuccal and lingual faces of the tooth, perpendicular to the MDdiameter. For molars, the BL diameter was measured as themaximum distance between the buccal and lingual surface ofthe tooth crown (Tobias, 1967). When the root was preserved, themaximum length from the cervical margin of the crown to the rootapex was measured.

Computer-based methods

High-resolution mCT images of the teeth were made at theVienna Micro-CT Lab, Department of Anthropology, University ofVienna, with a Viscom X8060 mCT scanner using the following scanparameters: 120 kV, 100 mA, voxel size¼ 20 mm. The resulting mCTdataset was first prepared for segmentation using a filteringprotocol developed at the Max Planck Institute for EvolutionaryAnthropology (Leipzig, Germany). This includes a Kiwahara anda 3Dmedian filter facilitating the separation of dentine and enamel.The filtered image stacks were segmented interactively in Amira 5.2(Mercury Computer Systems, Chelmsford, MA) to reconstructvirtual 3D models of the teeth. These models were used for variousanalyses, including: 1) description of the internal structure of theteeth (e.g., the EDJ), 2) quantification of the 2D cusp area and therelative cusp area (only for postcanine teeth), 3) quantification of3D dental tissue proportions and enamel thickness, and 4) OcclusalFingerprint Analysis (OFA).

Figure 2. Taddeo 1 (LC1): a) Labial view; b) Lingual view; c) Occlusal view; d) Distalview; e) Mesial view. The white bar in the figure is equivalent to 1 cm.

S. Benazzi et al. / Journal of Human Evolution 61 (2011) 377e387 379

Asapreliminary step for computer-based analyses (namely, the lastthree points mentioned above), the teeth were oriented in thefollowing way in Rapidform XOR2 (INUS Technology, Inc., Seoul,Korea): The cervical line of each tooth was manually digitized usinga curve and a best fit plane through the points of the curve wascomputed(cervicalplane). Thetoothwasthenrotateduntil thecervicalplane was parallel to the xy-plane of the Cartesian coordinate system.

To quantify 2D cusp areas in postcanine teeth, we digitallytraced the contour of the cusps along the primary fissures thatseparate them. This contour, together with the occlusal outline ofthe tooth, was orthogonally projected onto the cervical plane tomeasure the 2D cusp area and the relative cusp area (asa percentage of the total crown base area) (cf. Bailey, 2004).

To quantify 3D dental tissue proportions and enamel thickness,we followed the methods of Olejniczak (2006) and Olejniczak et al.(2008). Accordingly, we measured the enamel volume (mm3), thedentine volume (including also the volume of the crown pulpchamber e mm3), and the enamel-dentine junction (EDJ) surface(mm2). These measurements were used for the computation ofboth the average enamel thickness (AET¼ volume of enameldivided by the EDJ surface; index in millimeters) and the relativeenamel thickness index (RET¼AET divided by the cubic root ofdentine volume; scale-free index).

Occlusal Fingerprint Analysis (OFA, Kullmer et al., 2009) wasapplied to describe the spatial distribution of wear facet pattern ofthe teeth. Since wear facets are created by attritional occlusalcontacts between antagonistic teeth, they offer information aboutocclusal movements between upper and lower dentition (Kullmeret al., 2009). Moreover, they are particularly useful for matchingteeth and for their correct identification (Fiorenza et al., 2010).Detailed information about OFA is provided in previous contribu-tions (Ulhaas et al., 2004, 2007; Kullmer et al., 2009; Fiorenza et al.,2011) and is summarized in the Supplementary Online Materials.

Metric comparison

The MD and BL diameters of the four Taddeo teeth werecompared with those collected from western Eurasian Late Pleis-tocene human samples (see Supplementary Online Materials). Thesample included Neanderthals e N (Suzuki and Takai, 1970; Frayer,1978; Wolpoff, 1979; Trinkaus, 1983; Bailey and Hublin, 2006 andown measurements e BV), southwest Asian Middle PaleolithicHomo sapiens e MPHS (Frayer, 1978; Vandermeersch, 1981), Euro-pean Upper Paleolithic Homo sapiens e UPHS (Frayer, 1978;Shpakova, 2001; Hillson and Trinkaus, 2002; Teschler-Nicola,2006; and our own measurements e BV) and modern Homosapiens e MHS (our measurements e BV). The bivariate compari-sons and the 95% confidence ellipses were plotted using R software(version 2.11.1; R Development Core Team 2008). Comparative rootlength data for C1 and upper premolars (P3 and P4) were collectedfrom Bailey (2005) and Walker et al. (2008). The sample includedNeanderthals, UPHS and MHS. The comparative dataset for molarrelative cusp area was collected from Bailey (2002), and includesNeanderthals, MPHS, UPHS and MHS specimens. The comparativedataset for enamel thickness and dental tissue proportions wasobtained from Kono (2004) and Olejniczak et al. (2008). Unfortu-nately, this dataset includes only Neanderthal and modern humanmolars. A comparative sample for incisors, canines and premolars isnot available thus far.

Association between Taddeo 2 and Taddeo 3

To test the association between Taddeo 2 and Taddeo 3, bothdental surface models were imported into the Occlusal FingerprintAnalyzer (OFA) software. The OFA software is developed to move

virtually antagonistic tooth models towards each other alonga defined pathway, to simulate and analyze the collision of crowncontacts, e.g., during occlusion. OFA software prevents the pene-tration of the models thanks to collision detection, deflation andbreak free algorithms implemented into the application. Thetolerance distance for collision was set to 0.12 mm, slightly largerthan the average triangle length of the surface models. The trian-gles of the surfacemodels detected during contact between the twomodels were automatically selected and visualized in color.

Virtual reconstruction

A virtual reconstruction of the central region of the root wasrequired for Taddeo 1 and Taddeo 2, either to collect quantitativedental information or to correct previous restorations. In bothcases, the reconstructions were accomplished using 3D digitalmodeling and Computer-Aided Design (CAD) techniques. In detail,large lacunae were reconstructed by fitting NURBS (non-uniformrational basis-spline) surfaces across holes (Piegl Les, 1991; PieglLes and Tiller, 1996), while small holes were filled automaticallyin Rapidform XOR2. To evaluate the dentine volume of the root, weseparated the crown from the root based on the best fit planecomputed at the cervical line. For the canine, the mesial aspect ofthe cervical line was not considered in the computation of the bestfit plane, because the mesial aspect is too occlusally displacedcompared with the buccal, lingual and distal cervical line.

Results

Taddeo 1

Taddeo 1 is a complete lower left canine (LC1; Fig. 2). A largecrack traverses the crown in a disto-buccal to mesio-lingualdirection, and larger cracks also can be seen in the crown androot. From mCT images, it appears that the tooth had been recentlyreassembled from four pieces (Fig. 3a). Several smaller cracks arealso present, but do not seem to change the dimensions or theshape of the tooth significantly.

The tooth is heavily worn (category 5 of Molnar, 1971), resultingin considerable reduction of the crown height (4.3 mm) and analmost flat, distally sloping incisive surface with a large dentinebasin, enclosed by a residual enamel rim (see also theSupplementary Online Materials for detailed information aboutOFA results).

From the labial and lingual aspect, the mesial margin is almostvertical and the distal margin is slightly tilted distally. The labial

Figure 3. 3D digital model of Taddeo 1. a) On the left, the enamel cap (top) is separatedfrom the enamel-dentine junction (EDJ, bottom). The arrows point towards the mainfragments of the tooth. The best fit plane at the cervical line was computed withoutconsidering the mesial cervical aspect. b) The best fit plane was used to separate theroot from the crown. Missing portions of the root (in black) as well as the pulpchamber (right side of the picture) were reconstructed using CAD techniques, andvolumetric data were subsequently obtained.


surface is convex occluso-cervically and mesio-distally, with themaximum convexity in the lower third of the crown. The lingualsurface shows remnants of shoveling (ASUDAS grade 3). The distalmarginal ridge is marked and a distinct marginal groove follows itsedge. The mesial marginal ridge is less pronounced than the distalone and damaged by a vertical crack. The mesial and distal ridgesconverge in a slightly expressed basal eminence.

The distal side shows a well marked teardrop-shaped inter-proximal facet (2.9 mm� 1.8 mm), bucco-lingually extended(Fig. 3a), while the mesial one was removed by the occlusal wear.

Two linear enamel hypoplasias are visible in the buccal lowerthird of the crown, at 2.7 mm and 3.9 mm from the cervical line,respectively.

In its crown dimensions, the MD diameter is 7.5 mm and the BLdiameter is 8.2 mm. The root is 21 mm long, with a small crackseparating the apex from the rest of the root (Fig. 3a). It divergesslightly distally, particularly marked in the apical third. A broad andshallow longitudinal depression along the mesial root surface wasidentified in a former examination in 2001 by the first author.Currently, a figure-eight shaped hole about 5.4 mm (supero-infe-riorly) by 2.7 mm (bucco-lingually) made for aDNA sampling runsthrough the root in a mesio-distal direction, preventing theassessment of its original morphology. Additionally, probably inpreparation for the aDNA sampling, large parts of the surface werecarved away, as can be clearly seen in Fig. 2.

The virtual reconstructed volume of the removed root portion,carried out by CAD techniques, is 55.6 mm3 (Fig. 3b). The rootdentine volume, including the restored cracks and the portionremoved for aDNA sampling but without the pulp chamber, is366.54 mm3. This value underestimates the original root dentinevolume because of the carving away of the surface. Unfortunately,this missing portion of the dentine cannot be reconstructed atpresent. We also reconstructed the pulp chamber to quantify boththe root pulp volume (20.22 mm3; Fig. 3b) and the entire pulpvolume (root pulp volumeþ crown pulp volume¼ 26.31 mm3).

Metric comparison

The MD and BL diameters are at the upper limit of the MHSsample, but they fall near the average values of the other groups (N,

MPHS and UPHS) (Fig. 4a). In a scatter plot of the Neanderthal teeth,twomain groups can be recognized. Taddeo 1 is positioned near theNeanderthal group characterized by smaller diameters, similar insize to Vindija 288 (Fig. 4b). For root length, Taddeo 1 falls in theNeanderthal mean. It is larger comparedwith the UPHS sample andlies at the upper end of modern human variation (Table 1).

Taddeo 2

Taddeo 2 is a complete upper right fourth premolar (RP4; Fig. 5).Several small cracks are visible both externally and internally,crossing the enamel and dentine. On the external surface of thecrown, these cracks run along almost the entire surface from thecervix to the occlusal margin (Fig. 6a). The tooth is fairly worn(category 4e5 of Molnar, 1971), showing two small and separateddentine basins, developed on the buccal (paracone) and lingual(protocone) cusps (Fig. 5; see also the Supplementary OnlineMaterials for detailed information about OFA results).

The occlusal outline is oval, with an almost straight mesial sideand a convex distal one. The protocone (20.8 mm2, 46%) and theparacone (24.3 mm2, 54%) are subequal. The two cusps are con-nected by an almost completely worn mesial transverse crest,which marks the distal border of a mesial fossa. Distally, a star-shaped fossa can be observed, from which two barely visiblegrooves extend onto the distal marginal ridge, marking theremnants of a distal accessory marginal tubercle. The buccal face ofthe crown presents a small bulge in the lower third of the crown,which appears more marked due to the expression of a linearenamel hypoplasia along its occlusal limit, about 2.8 mm from thecervical line.

The mesial and distal interproximal facets are well marked, andquite wide bucco-lingually. The mesial facet is teardrop-shaped,with a supero-inferiorly very tall lingual half that gradually slopesout buccally, and measures 5.1 mm� 2.4 mm. The distal facet issubrectangular and measures 5.9 mm� 2.5 mm. In its crowndimensions, the MD diameter is 7.41 mm and the BL diameter is9.95 mm.

The EDJ shows a well-developed ridge connecting the paraconeand protocone horns (Fig. 6b), which would correspond with thetransverse crest identified in the occlusal surface. Furthermore,from each horn a further ridge bifurcates (more pronouncedlingually than buccally), suggesting that a bifurcate essential crestwas present on the occlusal surface. Distally, the EDJ surface is quitecomplex, marked by several accessory ridges. Both marginal ridgesare strongly developed. The distal one exhibits a small distalaccessory tubercle.

Two separate roots, one buccal and one lingual, originate froma long neck. The lingual root is quite straight, while the buccal root,broken between the middle and upper third, bends lingually(Fig. 6a). A hole about 3.3 mm in diameter made for aDNA samplingand subsequently filled with wax, pierces the root in a mesio-distaldirection. A further hole intersects the buccal root, probablycausing its fracture. Thereafter, the buccal root was incorrectlyglued to the tooth (Fig. 6a).

For this reason, the original morphology was reconstructed byvirtual means. Since part of the hole created by the drill used foraDNA sampling is still visible on the buccal root fragment (arrow inFig. 7a), we created the best fit cylinder of this surface (Fig. 7b). Thecylinder should approximately, but obviously not exactly, corre-spond to the drill bit. The buccal root fragment was interactivelydisplaced (in red in Fig. 7c) until the cylinder roughly matched theremnant of the hole in the other portion of the tooth. Finally, themissing parts were reconstructed using CAD techniques (Fig. 7d).Unfortunately, the pulp chamber was so extensively damaged bythe drill that its restoration was not feasible. The buccal root

Figure 4. Scatter plot between MD and BL diameters of the lower canine (a and b, the latter considering only Neanderthal), the P4 (c), the M1 (d), the M1 (e) and M2 (f).N¼Neanderthal, MPHS¼Middle Paleolithic Homo sapiens, UPHS¼ European Upper Paleolithic Homo sapiens, MHS¼modern Homo sapiens. In the scatter plots, the 95% confidenceellipses are reported for N, UPHS and MHS.


measures 14.5 mm and the lingual root is 14 mm. The volume of theroot was obtained by cutting the tooth with the best fit plane at thecervical line (root dentineþ root pulp¼ 311.31 mm3).

Metric comparison

Based on the MD and BL diameters, Taddeo 2 lies at the limit ofthe 95% confidence ellipse of the MHS, but falls well within therange of the other three groups (Fig. 4c). Like Taddeo 1, Taddeo 2

falls on the lower end of the Neanderthal size distribution. For rootlength, Taddeo 2 lies outside the lower range of the Neanderthal P4sand falls in the range of variation of MHS (Table 2).

Taddeo 3

Taddeo 3 is an upper right first molar (RM1) that has been gluedtogether from at least six pieces, some of which do not fit togetherperfectly. Several isolated fragments of dentine are also present.

Table 1Descriptive statistics of Neanderthal (N), Upper Paleolithic Homo sapiens (UPHS) andmodern Homo sapiens (MHS) C1 root lengths (mm) compared with Taddeo 1.

Tooth n Mean SD Range

N-C1a 8 19.4 2.4 16.3e23.2N-C1b 14 21.0 2.7 16.3e26.0UPHS-C1

a 6 16.2 2.3 13.1e19.0UPHS-C1

b 4 17.8 0.8 17.3e19.0MHS-C1 (Australian)a 18.1 17.0e21.0MHS-C1 (Bantu)a 17.0 12.5e20.5Taddeo 1 21.0

a Comparative data from Bailey (2005).b Comparative data from Walker et al. (2008).

Figure 6. 3D digital model of Taddeo 2. a) Mesial view of the tooth without the waxused for filling the holes. b) Enamel-dentine junction (EDJ) of Taddeo 2: a markedessential crest connects the buccal and the lingual horns. Its bifurcation is moreevident in the lingual horn (protocone). B¼ buccal; L¼ lingual; D¼ distal.


The crown is mostly complete, but the enamel from the mesio-buccal corner of the paracone is chipped away (Fig. 8). Two largecracks are visible, one on the buccal side of the crown, runningoccluso-cervically and continuing in the upper third of the root(Fig. 8a), and the second on the mesio-buccal corner of the para-cone, running distally on the occlusal surface parallel to the buccalside (Fig. 8d). The internal structure of the crown was partiallydamaged by aDNA sampling. Dentine inside the crown wasremoved by drilling starting from the pulp cavity.

The occlusal wear is strong (category 4e5 of Molnar, 1971), withfour large dentine exposures on the four main cusps, and a flatsurface in the trigon basin where the fissure pattern is almostcompletely obliterated (see the Supplementary OnlineMaterials fordetailed information about OFA results).

The occlusal outline is rhomboidal, mainly due to a lingualdisplacement of the metacone and a disto-lingual bulging of thehypocone (ASUDAS grade 4). The protocone is the largest cusp(28.1 mm2, 30.4%). The paracone (23.6 mm2, 25.5%; contour cor-rected following indications provided by Wood and Abbott, 1983;Bailey, 2002, 2004) and hypocone (22.2 mm2, 24%) are approxi-mately equal in size, while the metacone is the smallest one(18.5 mm2, 20%).

Figure 5. Taddeo 2 (RP4): a) Buccal view; b) Lingual view; c) Mesial view; d) Occlusalview; e) Distal view. The white bar in the figure is equivalent to 1 cm.

Both the mesial and distal interproximal facets are large andapparently subrectangular in shape, though the mesial facet isslightly damaged. The mesial interproximal facet measures4.8 mm� 2.5 mm,while the distal facetmeasures 6.2 mm� 2.7 mm.A thin linear enamel hypoplasia is visible at 1.2 mm from the cervicalline. In its crown dimensions, MD diameter is 11.2 mm and the BLdiameter is 12.2 mm. The two large cracks observed on the occlusalsurface are also visible in the EDJ (Fig. 9). Moreover, a third fractureruns mesio-lingually across the protocone. Despite the heavy wear,the EDJ still maintains important morphological features like a wellmarked oblique ridge, several accessory ridges on all the four cusps,strongly developed marginal ridges and a Carabelli’s tubercle. Thelatter feature shows that a Carabelli’s cusp was present in the mesio-lingual surface of the tooth. No other tubercles were observed on theEDJ, suggesting that a C5 (metaconule)was not present on the surfaceof the crown.

In 2001, when one of us (SB) studied this tooth, both buccal rootswere present (see also Messeri, 1975), while the lingual one wasfractured at the upper third. The two buccal roots diverged at4.8 mm from the cervical line (based on observations in 2001). Thedisto-buccal root is 11.8 mm long, while the mesio-buccal one is

Figure 7. Steps involved for virtual restoration of Taddeo 2’s root. a) The arrow pointsto the hole’s surface left by the drill, through which the best fit cylinder was computed(b). c) Distal view: the buccal root fragment was manually displaced as long as thecylinder approximately matches the surface of the upper hole’s surface left by the drill(the original fragment in gray, and the displaced portion in red). d) Distal view: finalreconstruction of the gaps (dark gray) using CAD techniques.

Table 2Descriptive statistics of Neanderthal (N), Upper Paleolithic Homo sapiens (UPHS) andmodern Homo sapiens (MHS) P4 root lengths (mm) compared with Taddeo 2.

Tooth n Mean SD Range

N-P4 10 17.6 1.0 16.2e19.0UPHS-P4 2 11.9 10.5e13.3MHS-P4 (Australian) 16.9 13.0e20.0MHS-P4 (Bantu) 14.6 13.0e19.5Taddeo 2 14.3a

Comparative data from Bailey (2005).a average between the buccal and the lingual root.

Figure 9. Enamel-dentine junction (EDJ) of Taddeo 3. B¼ buccal; L¼ lingual; M¼mesial.


13.2 mm. In its present state, the mesio-buccal root is brokenbetween themiddle and upper third, while for the disto-buccal rootonly the upper third is currently preserved.

Metric comparison

Both the MD and BL diameters are close to the averages acrossthe entire comparative sample (Fig. 4d). The pattern of relative cuspareas obtained for Taddeo 3 (mainly in relation to the hypocone)falls near the mean values computed for Neanderthal M1s (Table 3;see also Bailey, 2002). This pattern differs both from NeanderthalM2s and from the entire Homo sapiens comparative sample, exceptfor MPHS-M1s, which show a large hypocone and relativelyreduced metacone.

Taddeo 4

This tooth consists of a well preserved crown lacking the roots(Fig. 10). Several microfractures can be observed within the enameland dentine from mCT images. No dentine is exposed on the occlusalsurface, but small wear facets are visible (category 2 of Molnar,1971). The latter demonstrate that the tooth was in full functionalocclusion. Moreover, the presence of a large facet 12 on the disto-lingual slope of the hypoconulid, which is divided into two parts(a and b), points out a prominent hypocone in the upper molar (see

Figure 8. Taddeo 3 (RM1): a) Buccal view; b) Lingual view; c) Mesial view; d) Occlusalview; e). Distal view. The white bar in the figure is equivalent to 1 cm.

the Supplementary Online Materials for detailed information aboutOFA results).

As seen for Taddeo 3, the internal aspect of the crown wasirreversibly damaged by aDNA sampling, causing the removal ofa large portion of the crown dentine by drilling upwards from thepulp cavity (cf. Taddeo 3). The crown has a rectangular occlusaloutline. All five principal cusps are well-developed and arearranged in a Y-pattern. The protoconid is the largest cusp(34.2 mm2, 29.5%), followed by the metaconid (23.6 mm2, 20.4%).The hypoconid is well-developed (21 mm2, 18.1%), equals in size tothe entoconid (21 mm2, 18.1%). The hypoconulid is the smallestcusp (16 mm2,13.9%), though still well-developed (ASUDAS Cusp 5:grade 5). No C6 is present (Figs. 10 and 11).

Several extra fissures and crests are present on the main cusps.The mesial marginal ridge is quite thick. It limits the anterior foveainto a narrow and deep fissure (ASUDAS grade 4). An uninterruptedmid-trigonid crest (MTC), a continuous enamel ridge connectingthe protoconid and the metaconid, is well-developed, and it isabout the same height as the mesial marginal ridge. The medianridge of the metaconid is rather straight, without distal deflection.A distal trigonid crest was not observed. The distal marginal ridge isfairly thick and incised by a groove that continues onto the distalface separating the hypoconulid and entoconid (Fig. 10, 11). Themesial interproximal facet measures 4.2 mm� 2.3 mm, but there isno distal facet. A line of enamel hypoplasia that surrounds thewhole crown is present at 1.7 mm from the cervical line. In itscrown dimensions, MD diameter is 12.3 mm and the BL diameter is10.9 mm.

Table 3Relative cusp areas of Neanderthal (N), Middle Paleolithic Homo sapiens (MPHS),Upper Paleolithic Homo sapiens (UPHS), modern Homo sapiens (MHS) M1 and M2

compared with Taddeo 3.

Protocone Paracone Metacone Hypocone

Tooth n Mean SD Mean SD Mean SD Mean SD

N-M1 14 29.7 2.6 25.2 2.3 21.1 1.8 24.1 2.1N-M2 11 31.9 2.1 28.4 2.9 21.2 1.7 19.0 3.7MPHS-M1 3 29.5 3.1 23.5 0.8 19.6 0.1 27.4 2.9MPHS-M2 3 33.8 2.2 25.0 1.7 18.7 1.8 22.5 2.1UPHS-M1 6 30.7 1.7 25.9 2.3 23.5 2.2 20.0 4.0UPHS-M2 7 41.7 5.4 30.1 2.7 19.8 3.3 8.5 4.6MHS-M1 64 31.0 2.1 25.9 2.0 22.9 1.9 20.2 2.4MHS-M2 79 35.0 3.8 29.3 2.5 21.0 2.5 14.7 5.4Taddeo 3 30.4 25.5 20.0 24.0

Comparative data from Bailey (2002).

Figure 10. Taddeo 4 (RM1): a) Buccal view; b) Lingual view; c) Mesial view; d) Occlusalview; e) Distal view. The white bar in the figure is equivalent to 1 cm.


On the EDJ, the protoconid and metaconid are connected byamarked, mesially placedMTC, which carries a small tubercle closeto the median plane. Several accessory ridges are discernable,which generally mirror the external morphology of the enamelsurface. It is also worth noting the well marked bucco-distal groovethat separates the hypoconid and hypoconulid (Fig. 11).

Metric comparison

When compared with a sample of M1 and M2, the Taddeo 4diameters falls in the widely overlapping ranges of the four groups,except for its larger size compared with MHS M2s (Fig. 4e, f).

Considering the relative cusp areas, a large protoconid followedby a metaconid larger or similar in size to the entoconid is quitenormal in M1s, both in Neanderthals and in UPHS-MHS (Table 4).Beside this, the extremely large protoconid obtained from Taddeo 4is outside the range of variation computed for Neanderthal M1 thusfar.

The Taddeo 4 crown is characterized by large enamel volumeand low “dentine þ pulp” volume (Table 5), which together causehigh Average and Relative Enamel thickness indices(AET¼ 1.32 mm; RET¼ 19.67). Taddeo 4 falls at the high end of theNeanderthal M2 range of variation in enamel thickness, surpassingall nine Neanderthal M1s published by Olejniczak et al. (2008)(Table 5). Conversely, Taddeo 4 dentine and pulp volume is in thelower range of the Neanderthal variation, both for M1 and M2. Inthese features, Taddeo 4 is more similar to modern humans than toNeanderthals as they have been described to date. The AET indexcomputed for Taddeo 4 matches the average value provided byKono (2004) for modern humanM1s, but it falls in the high range ofvariation for Neanderthal M1s (outside the range computed for theNeanderthal M2s). Both for Neanderthal and modern human, Tad-deo 4 is outside the range of variation measured for M3s byOlejniczak et al.(2008; data for M3s are not shown in Table 5).

Figure 11. Taddeo 4: 3D digital model of the reconstructed crown (left) and enamel-dentine junction (EDJ, right). B¼ buccal; L¼ lingual; M¼mesial.

Association between Taddeo 2 and Taddeo 3

As shown in Fig. 12a, the teeth were considered to be correctlypositioned when both the continuity of their occlusal surfaces wasrestored and the interproximal facets were in contact.

In this condition, the mesial interproximal facet of Taddeo 3(Fig. 12b) perfectly matches the distal interproximal facet of Taddeo

2 (Fig. 12c). A lack of contact on the buccal aspect of the latter is dueto the fracture of the paracone of Taddeo 3 mentioned above (seearrow in Fig. 12c). Additionally, results from Taddeo 2 and 3 wearfacet pattern analysis (see Supplementary Online Materials) showthat wear patterns of both specimens are in a similar wear stage.The Occlusal Fingerprint Analysis strongly supports the associationbetween the specimens.

Discussion

Classification

Two different identifications were proposed for Taddeo 1: firstan upper right canine (based on the small size of the tooth and thepresence of a lingual basal eminence; Messeri, 1975), and thena lower right canine by one of the co-authors (FM, personalcommunication in Giacobini and Manzi, 2005). We disagree withboth and identify Taddeo 1 as a lower left canine (LC1). The shape ofthe crown, with relatively slight flaring distally and mesially, and ingeneral a slender aspect is a much better fit with lower than uppercanines. The shape of the marginal ridges and the absence ofa marked lingual buttress also support this attribution. Siding isbased on the curvature of the root tip, the strongly distally slopingocclusal wear, as well as the occlusal outline.

Taddeo 2 was previously classified as a RP3 based on the largebuccal cusp, the straight (not curved) sulcus that splits the twocusps, and the bifurcated root (Messeri, 1975; Messeri and Palma diCesnola, 1976). Nevertheless, based on several features this toothshould be considered a RP4. These features include in particular theoval shape of the occlusal outline, which is different from the moretriangular shape seen in P3s, the small size difference betweenbuccal (paracone) and lingual (protocone) cusp, and the largebucco-lingually extended mesial interproximal facet (which couldhardly be created by canine contact). Moreover, the bifurcateessential crest and especially the bifurcation in the lingual horndetected in the EDJ of Taddeo 2 has been observed more frequentlyin P4 (70%) than in P3 (58%) occlusal surfaces (Bailey, 2006), furthersupporting its classification to P4. Other morphological featuresrecognized in the EDJ (e.g., the large number of accessory ridges andthe distal accessory marginal tubercle), are not informative forpurposes of identification because they appear in similar frequencyin both upper premolars (Bailey, 2002).

We confirm the previous classification for Taddeo 3 mainly withregard to its pattern of relative cusp areas rather than on itsmorphological traits. In fact, it has been observed that morpho-logical features like the frequency for Carabelli’s cusp, C5 andmesial accessory tubercles, are as likely to occur on theM2 as on theM1 (Bailey, 2002). Moreover, the association between Taddeo 2 andTaddeo 3 has been established based on the similarity in wear

Table 4Relative cusp areas of Neanderthal (N), Middle Paleolithic Homo sapiens (MPHS), Upper Paleolithic Homo sapiens (UPHS), modern Homo sapiens (MHS) M1 and M2 comparedwith Taddeo 4.

Protoconid Metaconid Hypoconid Entoconid Hypoconulid

Tooth n Mean SD Mean SD Mean SD Mean SD Mean SD

N-M1 12 24.2 2.2 23.6 3.2 19.3 3.9 20.8 3.8 12.1 3.1N-M2 14a 26.1 2.9 21.3 3.8 23.2 4.1 20.3 2.8 10.1 4.4MPHS-M1 2 23.6 28.0 18.0 16.9 12.4MPHS-M2 2 24.3 27.5 19.2 21.1 8.0UPHS-M1 4b 26.3 2.7 22.1 1.2 22.0 3.4 20.3 2.6 9.3 6.6UPHS-M2 7c 29.8 2.1 21.4 3.5 26.3 3.0 21.5 3.2 7.1MHS-M1 63d 24.9 2.1 22.3 1.9 20.1 2.4 20.6 2.0 12.1 3.8MHS-M2 88e 26.5 2.8 22.8 3.1 24.7 4.2 22.2 3.7 10.4 3.7Taddeo 4 29.5 20.4 18.1 18.1 13.9

Comparative data from Bailey (2002).a 12 Specimens for the hypoconulid.b Three specimens for the hypoconulid.c One specimen for the hypoconulid.d 61 Specimens for the hypoconulid.e 33 Specimens for the hypoconulid.


pattern between the RP4 and the RM1, and the perfect matchbetween the distal interproximal facet of Taddeo 2 and the mesialinterproximal facet of Taddeo 3.

Several features support the identification of Taddeo 4 as an M1instead of an M2, confirming the previous classification provided byMesseri (1975): 1) the regular morphology of this molar, 2) thepattern of relative cusp areas (metaconid> entoconid), 3) theabsence of C6 (this cusp is more frequent in M2 (50%) than on M1(36%); Bailey, 2006), and 4) the well-developed facets 12a and 12b(pointing out a prominent hypocone in the upper molar).

Taxonomic assessment

Although Taddeo 1 is heavily worn, a good analogue for themorphological features of this tooth crown (but not the root) is theleft canine of Krapina 138, lending further support to our identifi-cation (see Supplementary Online Materials). In the lingual aspect,both specimens show a remnant of shoveling, a marked distalmarginal ridge with a distinct marginal groove delimiting its edge,and a slightly expressed basal eminence, in which the mesial anddistal ridges converge. Taddeo 1 is also similar to Vindija 288 in itssmall crown size, but falls near the Neanderthal average for rootlength. A small crown size could represent a peculiarity ofa southern Neanderthal group, as suggested previously (de Lumley,1973). Alternatively, the canine of Taddeo cave belonged toa Neanderthal female. However, at present we lack the necessaryinformation about the variability related to sexual dimorphism inNeanderthals to evaluate this hypothesis.

The bifurcated essential crest and the distal accessory marginaltubercle visible in Taddeo 2 support its Neanderthal classification.Bailey (2006) observed that the morphological variation of the

Table 5Enamel thickness of Taddeo 4 compared with Neanderthal (N) and modern Homo sapien

Tooth Taxon n Enamel volume(mm3)

Coronal dentineþ pvolume (mm3)

M1 Na 9 232.73 (18.58) 345.45 (54.72)MHSa 1 215.65 289.83MHSb 13 279.40 (37.70) 304.60 (47.30)

M2 Na 4 232.95 (72.71) 391.02 (102.12)MHSa 9 235.67 (49.75) 245.61 (60.65)MHSb 13 252.40 (36.10) 272.10 (59.00)

Taddeo 4 290.78 301.85

a Data from Olejniczak et al. (2008).b Data from Kono (2004).

Neanderthal P4s generally falls in the range of variation of MHS,except for the higher frequency of a bifurcated essential crest andthe distal accessory cusp. The latter features slightly distinguishNeanderthal P4 even in comparison with other fossil humans(Bailey, 2006). Finally, it is worth noting that our morphometricanalysis of the crown diameters and root length of this tooth placesit in the lower end (or even outside for the root length) of theNeanderthal range of variation. As with the canine, this couldindicate sexual or regional patterns in the Neanderthal variation.

The variability of both canine root length and P4 root lengthseems to be underestimated in the data given by Bailey (Adeline LeCabec, pers. comm, BV, unpublished data). Accordingly, we cautionagainst the use of root length as a method for taxonomic assess-ment until further work has been completed on this subject.

Unfortunately, we do not have a Neanderthal and modernhuman comparative dataset for enamel thickness and dental tissueproportions of canines and premolars. Due to the heavy wear ofTaddeo 1 and Taddeo 2, an evaluation of the original enamelthickness would not be feasible. Even so, we have provided data onroot dentine (for Taddeo 1 and Taddeo 2) and pulp chamber (forTaddeo 1) that will be useful for future comparative work.

Based on the pattern of relative cusp areas (large hypocone andrelative small metacone), Taddeo 3 falls in the mean valuescomputed for Neanderthal M1s (data from Bailey, 2002; see alsoBailey, 2004; Quam et al., 2009).

There are two important morphological features supporting theNeanderthal classification of Taddeo 4 e namely, an anterior fovea,which is distally bordered by a well marked mid-trigonid crest (DeLumley, 1973; Bailey, 2002, 2006). It has been demonstrated thatthe combination of an anterior fovea and a continuousmid-trigonidcrest is a relevant morphological character to distinguish between

s (MHS) M1s and M2s (SD in brackets).

ulp EDJ surface area(mm2)

Average enamelthickness (mm)

Relative enamelthickness (scale-free)

207.37 (46.75) 1.17 (0.23) 16.77 (3.8)205.32 1.05 15.87211.90 (23.70) 1.32 (0.12)231.56 (52.18) 0.99 (0.10) 13.63 (0.47)177.79 (61.58) 1.46 (0.49) 23.44 (8.35)194.80 (27.10) 1.30 (0.13)220.42 1.32 19.67

Figure 12. a) Virtual matching of Taddeo 2 (RP4) and Taddeo 3 (RM1) applying the OFAsoftware. b) Taddeo 2 is transparent to show the collision in the mesial interproximalfacet of Taddeo 3. c) Taddeo 3 is transparent to show the collision area detected in thedistal interproximal facet of Taddeo 2. The arrow points towards the area of theinterproximal facet where no collision was detected by the OFA software.


Neanderthal and early modern Europeans (Bailey, 2006). Results ofthe quantification of the enamel thickness and tissue proportionsare more ambiguous. Previous analyses based on two-dimensionaldata suggest that Neanderthal molars have in general thinnerenamel than modern humans (Zilberman et al., 1992, Zilbermanand Smith, 1992). More recently, Olejniczak et al. (2008) pre-sented detailed 3D data on Neanderthal enamel thickness, showingthat it is both absolutely and relatively (i.e., relative to dentinevolume) thinner than in modern humans. Nonetheless, the Taddeo4 crown shows the largest enamel thickness and among the lowest“dentineþ pulp” volume of every Neanderthal M1 (n¼ nine) pub-lished by Olejniczak et al. (2008). In its absolute dimensions, as wellas for the AET index, Taddeo 4 resembles modern humanM1s morethan it does Neanderthal M1s.

We believe that Taddeo 4 shows the limits of the enamelthickness volume and dental tissue proportions for taxonomicpurposes, suggesting caution in using this technique as a standalone discriminatory parameter until the Neanderthal range ofvariation is better understood.

Based on the advanced wear stage of LC, RP4 and RM1, and thealmost unworn condition of the lower molar, the four teethdiscovered in Taddeo cave belong at least to two individuals. On thebasis of the wear stage, the lower molar does not fit with the otherteeth. Even if it were an M3, it should show at least wear stage 3 tomatch the wear pattern of the other teeth. Accordingly, the lowermolar belongs to a second, younger individual. However, we arecertain of the relationship between the two upper teeth, whicharticulate well together as it is evident through the OcclusalFingerprint Analyser test result. On the basis of a similar stage ofocclusal wear, and the small crown size similar to the RP4, the lowercanine could also be attributed to the same individual.

Conclusions

The human teeth discovered in Taddeo cave (Marina di Camer-ota coast, Campania, Italy) have been considered to representNeanderthals, despite the lack of detailed morphological andmetric analyses. In the present contribution, we have provideda reassessment of these human teeth. We have rectified themisidentifications of Taddeo 1 (LC1) and Taddeo 2 (RP4), and haveestablished the association between Taddeo 2 and Taddeo 3, basedon the fit of the interproximal facets and the continuity of the wearpatterns. We agree with the previous morphological diagnosis ofTaddeo 4 (RM1).

All of the teeth show clear Neanderthal morphological features.For Taddeo 3, it is confirmed by the pattern of relative cusp areas.Regarding Taddeo 4, we also point out that the enamel thicknessand dental tissue proportions resemble modern humans more thanNeanderthal M1s, thus increasing the known range of variation ofNeanderthals. This large range of variation is further supported bythe small size of Taddeo 1 and Taddeo 2, which could representa peculiarity of a southern Neanderthal group or could be explainedby sexual dimorphism.

Acknowledgments

The authors thank S. Kozakowski and B. Metscher for copy-editing of our manuscript. We are grateful to D. Byer for helpwith the English. We thank M. Skinner and S. Bailey for discussionsregarding the EDJ morphology of the Taddeo specimens. Finally,we would like to thank R. Ginner for taking photographs andA. Stadlmayr for casting the teeth from Taddeo cave. This researchwas supported by the “Deutsche Forschungsgemeinschaft” (DFG,German Research Foundation) and is publication no. 30 of the DFGResearch Unit 771 “Function and performance enhancement in themammalian dentition e phylogenetic and ontogenetic impact onthe masticatory apparatus.” This work was supported by EU MarieCurie Training Network MRTN-CT-2005-019564 EVAN and NSF 01-120 Hominid Grant 2007.

Appendix. Supplementary data

Supplementary data associated with this article can be found inthe online version, at doi:10.1016/j.jhevol.2011.05.001.

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A reassessment of the Neanderthal teeth from Taddeo cave (southern Italy)